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Unmanned Aerial Vehicles (UAVs), or drones, offer the ability to collect cost-effective fine-scale imagery that is suitable for the capture of concurrent hydrodynamic and faunal data within tidal stream environments. This is a necessary stage of information gathering to inform tidal energy device design, advise control and maintenance strategies and better inform environmental consenting processes. For this study a total of sixty-three UAV surveys were undertaken within the Inner Sound of the Pentland Firth, Scotland, UK, over two 4-day periods in 2016 and 2018. The aims of this data collection effort were to characterise bathymetrically driven hydrodynamic features, comprising of kolk-boil distribution, presence, and area, as well as marine life such as seabird distributions, presence, and orientation relative to the flow. To achieve this, a method to extract quantifiable metrics from UAV imagery was required. This paper details the processes and methodology to create a graphical user interface (GUI) to provide these outputs rather than examining specific results. It includes an explanation of the criteria that the GUI needed to meet to be able to process the imagery, a description of the workflow and an explanation of the sub-routines required such as image registration and calibration. The outputs of the GUI, and their relevance to tidal energy developments, are also discussed. Finally, this paper details future work incorporating computer vision techniques to improve the accuracy, reliability, and processing speed of the GUI.

Understanding spatiotemporally varying animal distributions can inform ecological understanding of species' behavior (e.g., foraging and predator/prey interactions) and support development of management and conservation measures. Data from an array of echolocation‐click detectors (C‐PODs) were analyzed using Bayesian spatiotemporal modeling to investigate spatial and temporal variation in occurrence and foraging activity of harbor porpoises (Phocoena phocoena) and how this variation was influenced by daylight and presence of bottlenose dolphins (Tursiops truncatus). The probability of occurrence of porpoises was highest on an offshore sandbank, where the proportion of detections with foraging clicks was relatively low. The porpoises' overall distribution shifted throughout the summer and autumn, likely influenced by seasonal prey availability. Probability of porpoise occurrence was lowest in areas close to the coast, where dolphin detections were highest and declined prior to dolphin detection, leading potentially to avoidance of spatiotemporal overlap between porpoises and dolphins. Increased understanding of porpoises' seasonal distribution, key foraging areas, and their relationship with competitors can shed light on management options and potential interactions with offshore industries.

Primary production dynamics are strongly associated with vertical density profiles, which dictate the depth of stratification and mixed layers. Climate change and artificial structures (e.g. windfarms) are likely to modify the strength of stratification and vertical distribution of nutrient fluxes, especially in shelf seas where fine scale processes are important drivers, affecting the vertical distribution of phytoplankton. To understand the effect of physical changes on primary production, identifying the linkage between density and phytoplankton profiles is essential. Here, the ecological relevance of eight density layers (DLs) obtained by multiple methods that define three different portions of the pycnocline (above, centre, below) was evaluated to identify a valuable proxy for subsurface Chlorophyll-a (Chl-a mg m-3) concentrations. The associations of subsurface Chl-a with surface and deep mixing were investigated by hypothesizing the occurrence at the same depth of any DL and the maximum Chl-a layer (DMC) using Spearman correlation, linear regression, and a Major Axis analysis. Out of 1237 observations of the water column exhibiting a pycnocline, 78 % reported DMCs above the bottom mixed layer depth (BMLD). This suggests that the BMLD is a boundary trapping Chl-a in shallow waters (≤ 120 m). BMLD constantly described Chl-a vertical distribution despite surface mixing indicators, suggesting a significant contribution of deep mixing processes in supporting subsurface production under specific conditions (e.g. prolonged stratification, tidal cycle, and bathymetry). Using BMLD for defining subsurface Chl-a could be a valuable tool for understanding the spatiotemporal variability of Chl-a in shelf seas, representing a potential variable for ecological assessments.

Changes in animal movement and behaviour at fine scales (tens of metres) in immediate proximity to tidal stream turbine structures are largely unknown and have implications for risks of animal collision with turbine blades. This study used upward-facing multibeam echosounder data to detect and track animal movement comprising fish, diving seabirds, and marine mammals. Measurements over spring-neap tidal cycles at a turbine structure (no blades present) are compared to a neighbouring reference area with no structure and comparable conditions, with measurements consecutive in time to maximize comparability.The majority of tracked animals (93.4% around turbine structure and 99.1% without turbine structure) were observed swimming against the flow, with 87.5% and 97.8%, respectively, making ground and showing capability of manoeuvring in tidal stream flow speeds. Track tortuosity increased around the turbine structure compared to the reference site, particularly in the wake and at low flow speeds, indicating animal station-holding or milling behaviour. These data also evidence the benefits of multibeam echosounders to measure animal movement through larger measurement volumes rather than relying on single-beam echosounders to measure animal presence alone, including to avoid large biases overestimating the size of schools swimming against the flow measured by time-in-beam.

Identifying and quantifying the effects of climate change that alter the habitat overlap of marine predators and their prey population distributions is of great importance for the sustainable management of populations. This study uses Bayesian joint models with integrated nested Laplace approximation (INLA) to predict future spatial density distributions in the form of common spatial trends of predator–prey overlap in 2050 under the “business‐as‐usual, worst‐case” climate change scenario. This was done for combinations of six mobile marine predator species (gray seal, harbor seal, harbor porpoise, common guillemot, black‐legged kittiwake, and northern gannet) and two of their common prey species (herring and sandeels). A range of five explanatory variables that cover both physical and biological aspects of critical marine habitat were used as follows: bottom temperature, stratification, depth‐averaged speed, net primary production, and maximum subsurface chlorophyll. Four different methods were explored to quantify relative ecological cost/benefits of climate change to the common spatial trends of predator–prey density distributions. All but one future joint model showed significant decreases in overall spatial percentage change. The most dramatic loss in predator–prey population overlap was shown by harbor seals with large declines in the common spatial trend for both prey species. On the positive side, both gannets and guillemots are projected to have localized regions with increased overlap with sandeels. Most joint predator–prey models showed large changes in centroid location, however the direction of change in centroids was not simply northwards, but mostly ranged from northwest to northeast. This approach can be very useful in informing the design of spatial management policies under climate change by using the potential differences in ecological costs to weigh up the trade‐offs in decisions involving issues of large‐scale spatial use of our oceans, such as marine protected areas, commercial fishing, and large‐scale marine renewable developments., This study tests a range of analytical techniques to quantify ecological costs of climate change on habitat availability of predator–prey or competing mobile species. The study demonstrates how to detect environmentally sensitive habitat areas and is highly useful for habitat management policies under climate change.

There is uncertainty on the ecological effects of tidal stream turbines. Concerns include animal collision with turbine blades, disruption of migratory and foraging behaviour, attraction of animals to prey aggregating around turbines, or conversely displacement of animals from preferred habitat. This study used concurrent ecological and physical measurements to show the predictability of fish school characteristics (presence, school area and height above seabed) in a high energy tidal site across spring/neap, ebb/flood and daily cycles, and how this changed around a turbine structure. The rate of schools and school area per hour increased by 1.74 and 1.75 times respectively around a turbine structure compared to observations under similar conditions without a turbine structure. The largest schools occurred at peak flow speeds and the vertical distribution of schools over the diel cycle was altered around the turbine structure. While predictable attraction or aggregation of prey may increase prey availability and predator foraging efficiency, attraction of predators has the potential to increase animal collision risk. Predictable changes from the installation of turbine structures can be used to estimate cumulative effects on predators at a population level. This study can guide a strategic approach to the monitoring and management of turbines and arrays.

D.C., J.M., P.T. and S.W. acknowledge the financial support of the Tidal Stream Industry Energiser project (TIGER), which is co-financed by the European Regional Development Fund through the Interreg France (Channel) England Programme. A.A. acknowledges the support of NERC through the Industrial Innovation fellowship grant no. NE/R013209/2. D.G., B.S., T.S., D.W., R.W. and B.W. acknowledge EPSRC for supporting part of this work through EP/S000747/1. G.H. and C.S. acknowledge support by the Scottish government-funded Marine Mammal Scientific Support Programme MMSS/002 with additional resources from the Natural Environment Research Council grant nos. NE/R014639/1 and NE/R015007/1. M.L. acknowledges the support of the Smart Efficient Energy Centre (SEEC) at Bangor University, part-funded by the European Regional Development Fund (ERDF), administered by the Welsh Government, and the EPSRC METRIC fellowship grant no. (EP/R034664/1). L.M. acknowledges the financial support of an EPSRC PhD studentship award, grant no. EP/R513052/1(18-22). J.McN. acknowledges the financial support of R.Willden’s EPSRC Advanced Fellowship grant no. EP/R007322/1. S.N. acknowledges the support of SEEC (Smart Efficient Energy Centre) at Bangor University, part-funded by the European Regional Development Fund (ERDF), administered by the Welsh Government. This review provides a critical, multi-faceted assessment of the practical contribution tidal stream energy can make to the UK and British Channel Islands future energy mix. Evidence is presented that broadly supports the latest national-scale practical resource estimate, of 34 TWh/year, equivalent to 11% of the UK’s current annual electricity demand. The size of the practical resource depends in part on the economic competitiveness of projects. In the UK, 124 MW of prospective tidal stream capacity is currently eligible to bid for subsidy support (MeyGen 1C, 80 MW; PTEC, 30 MW; and Morlais, 14 MW). It is estimated that the installation of this 124 MW would serve to drive down the levelized cost of energy (LCoE), through learning, from its current level of around 240 £/MWh to below 150 £/MWh, based on a mid-range technology learning rate of 17%. Doing so would make tidal stream cost competitive with technologies such as combined cycle gas turbines, biomass and anaerobic digestion. Installing this 124 MW by 2031 would put tidal stream on a trajectory to install the estimated 11.5 GW needed to generate 34 TWh/year by 2050. The cyclic, predictable nature of tidal stream power shows potential to provide additional, whole-system cost benefits. These include reductions in balancing expenditure that are not considered in conventional LCoE estimates. The practical resource is also dependent on environmental constraints. To date, no collisions between animals and turbines have been detected, and only small changes in habitat have been measured. The impacts of large arrays on stratification and predator–prey interaction are projected to be an order of magnitude less than those from climate change, highlighting opportunities for risk retirement. Ongoing field measurements will be important as arrays scale up, given the uncertainty in some environmental and ecological impact models. Based on the findings presented in this review, we recommend that an updated national-scale practical resource study is undertaken that implements high-fidelity, site-specific modelling, with improved model validation from the wide range of field measurements that are now available from the major sites. Quantifying the sensitivity of the practical resource to constraints will be important to establish opportunities for constraint retirement. Quantification of whole-system benefits is necessary to fully understand the value of tidal stream in the energy system. Publisher PDF

In the dynamic environments targeted for marine renewable energy extraction, such as tidal channels, the natural distribution of fish and behavioural impacts of marine renewable energy installations (MREIs) are poorly understood. This study builds on recent methodological developments to reveal the behaviour of fish schools using data collected by a seabed-mounted echosounder deployed in extreme tidal flows and in the wake of a MREI (composed of the foundation of a full-scale tidal stream energy turbine). The results show a significant change (p

The aim of this work is to analyse the potential impacts of tidal energy extraction on the marine environment. We wanted to put them in the broader context of the possibly greater and global ecological threat of climate change. Here, we present how very large (hypothetical) tidal stream arrays and a ''business as usual'' future climate scenario can change the hydrodynamics of a seasonally stratified shelf sea, and consequently modify ecosystem habitats and animals’ behaviour.The Scottish Shelf Model, an unstructured grid three-dimensional ocean model, has been used to reproduce the present and the future state of the NW European continental shelf. While the marine biogeochemical model ERSEM (European Regional Seas Ecosystem Model) has been used to describe the corresponding biogeochemical conditions. Four scenarios have been modelled: present conditions and projected future climate in 2050, each with and without very large scale tidal stream arrays in Scottish Waters (UK). This allows us to evaluate the potential effect of climate change and large scale energy extraction on the hydrodynamics and biogeochemistry. We found that climate change and tidal energy extraction both act in the same direction, in terms of increasing stratification due to warming and reduced mixing, however, the effect of climate change is ten times larger. Additionally, the ecological costs and benefits of these contrasting pressures on mobile predator and prey marine species are evaluated using ecological statistical models.

This work was funded by the Scottish Government project SP004 and a MSS PhD studentship to DS. Many thanks are due to the Crews of the MV Alba na Mara (MSS), RV Sir John Murray (SEPA) and the MV Lochnevis (Caledonian Macbrayne) for facilitating sample collection, John Dunn for assistance with the manufacture and installation of the ferry sampler, Marian Thomson and other staff at the University of Edinburgh for laboratory assistance, Anastasia Imsiridou, Sofia Galinou-Mitsoudi and Vassilis Katsares of the Greek Department of Fisheries and Aquaculture Technology for supplying reference adult A. fragilis DNA, Pablo Diaz and staff at the University of Aberdeen microscopy department for assistance with SEM analysis, the National Museum of Wales for allowing reproduction of the juvenile A. fragilis image, Keith Hiscock and Eve Southward of Plymouth Marine Laboratory for historical information on the identification of A. fragilis larvae, Colin McAlister and the staff of the Fishery offices in Mallaig and Fraserburgh for assistance in the transport of zooplankton samples and materials, and the British Oceanographic Data Centre for supplying data on the UK Tidal Gauge Network. Comments from Associate Editor Simon Cragg and two anonymous reviewers were greatly appreciated for improving the manuscript

Understanding ecosystem dynamics within shallow shelf seas is of great importance to support marine spatial management of natural populations and activities such as fishing and offshore renewable energy production to combat climate change. Given the possibility of future changes, a baseline is needed to predict ecosystems responses to such changes. This study uses Bayesian techniques to find the data-driven estimates of interactions among a set of physical and biological variables and a human pressure within the last 30 years in a well-studied shallow sea (North Sea, UK) with four contrasting regions and their associated ecosystems. A hidden variable is incorporated to model functional ecosystem change, where the underlying interactions dramatically change, following natural or anthropogenic disturbance. Data-driven estimates of interactions were identified, highlighting physical (e.g. bottom temperature, potential energy anomaly) and biological variables (e.g. sandeel larvae, net primary production) to be strong indicators of ecosystem change. There was consistency in the physical and biological variables, identified as good indicators in three of the regions, however the shallower region (with depths

High-flow tidal stream environments, targeted for tidal turbine installations, exhibit turbulent features, at fine spatio-temporal scales (metres and seconds), created by site-specific topography and bathymetry. Bed-derived turbulent features (kolk-boils) are thought to have detrimental effects on tidal turbines. Characterisation of kolk-boils is therefore essential to inform turbine reliability, control, and maintenance strategies. It will also improve the understanding of potential ecological interactions with turbines, as marine animals use these sites for foraging. Unmanned aerial vehicle (UAV), or drone, imagery offers a novel approach to take precise measurements of kolk-boil characteristics (distribution, presence, and area) at the surface. This study carried out sixty-three UAV surveys within the Inner Sound of the Pentland Firth, Scotland, UK, over four-day periods in 2016 and 2018. Kolk-boil characteristics were examined against relevant environmental covariates to investigate potential drivers of presence and area. The results show that distribution at the surface could be predicted based on tidal phase, with current velocity significantly influencing presence above 3.0 m/s. The technique can be used to inform turbine development, micro-siting and provide better understanding of environmental implications of turbine operation. Finally, it highlights the suitability of UAVs for capturing rapid fine-scale hydrodynamic data in the absence of in situ measurements.

Field measurements of the flow in the benthic boundary layer (BBL) of a tidal channel are presented which compare data collected in the wake of a marine renewable energy installation (MREI) with control data representative of the natural conditions. The results show significant flow modification in the wake of the MREI including a reduction in mean velocity, enhanced turbulence, and the breakdown of the natural structure and dynamics of the BBL. This study provides new information relevant to the environmental impact assessment of MREIs and to the design and consenting of marine renewable energy projects.

As harbour porpoises Phocoena phocoena are abundant within tidal stream environments, mitigating population-level impacts from tidal stream energy extraction is considered a conservation priority. An understanding of their spatial and temporal occupancy of these habitats at a regional-scale will help steer installations towards locations which maximize energy returns but reduce the potential for interactions with populations. This study quantifies and compares relationships between the presence of harbour porpoise and several hydrodynamic characteristics across four tidal stream environments in Anglesey, UK—a region that has been earmarked for extensive industrial development. Within sites (0.57–1.13 km2), encounters with animals were concentrated in small areas (2.5 ms−1), and placing turbines in areas of laminar currents therein. This study shows that a combination of local and regional hydrodynamic characteristics can partially explain variations in occupancy patterns across tidal-stream environments. However, it was found that such hydrodynamic characteristics could not comprehensively explain these occupancy patterns. Further studies into the biophysical mechanisms creating foraging opportunities within these habitats are needed to identify alternative explanatory variables that may have universal applications.

The global increase in tidal stream turbine installations creates a need to identify and mitigate any impacts on seabird populations. Within Scotland, UK, the vulnerability of black guillemots Cepphus grylle and European shags Phalacrocorax aristotelis is dependent on their tendency to exploit microhabitats characterised by fast mean horizontal current speeds (≥2 ms –1 ), and tidal states with maximum current speeds, within tidal stream environments. Identifying consistencies in their relative use of different microhabitats (fast versus slow mean horizontal current speeds) and tidal states (increasing/decreasing versus maximum currents) across these habitats could assist risk assessment and mitigation measures at both a regional and development site level. Datasets from shore-based surveys collated across 6 tidal stream environments showed that the probability of detecting foraging black guillemots and European shags tended to be higher in fast and slow microhabitats, respectively. However, differences between microhabitats were reversed and/or marginal in 3 out of the 5 sites used for each species. Differences between tidal states were almost always marginal. These variabilities show that a species' vulnerability could differ greatly among development sites, and environmental impact assessments (EIA) must quantify habitat-use using dedicated and site-specific surveys to reduce uncertainty. However, a greater understanding of the mechanisms underlying variation in the use of tidal stream environments is needed when selecting a suite of potential development sites that reduce the possibility of population-level impacts. The current collection of physical and biological data across tidal stream environments could therefore prove invaluable for the protection of seabird populations.

A detailed understanding of fishing activity in Scottish waters is required to inform marine spatial planning. Larger fishing vessels are fitted with Vessel Monitoring Systems (VMS) offering spatial information on fishing activity. VMS does not cover smaller vessels (under 15 m), which fish predominantly in inshore waters where the competition for space is often greatest. To improve knowledge of the distribution of fishing activity and value of fisheries in Scotland's inshore waters, Marine Scotland conducted a participatory fisheries mapping project, known as ScotMap. The data were collected during face-to-face interviews with 1090 fishermen of Scottish registered commercial fishing vessels under 15 m in overall length and relate to fishing activity for the period 2007–2011. Interviewees were asked to identify the areas in which they fish, estimate the contribution these areas make to vessel earnings, and to provide associated information. The majority of interviews relate to creel fishing. The data collected were aggregated to provide mapped outputs of the monetary value, relative importance to fishermen and the usage of the seas around Scotland (number of fishing vessels and number of crew). ScotMap outputs provide information on the locations of inshore fishing activities and the economic importance of different sea areas at a much higher spatial resolution than was previously possible. Outputs have informed marine policy development, provide a valuable resource for marine spatial planning in Scotland and illustrate how participatory mapping can generate useful resources on the location and importance of inshore fishing areas.

Funded by University of Aberdeen Natural Environment Research Council (NERC) Department for Environment, Food, and Rural Affairs. Grant Number: NE/J004308/1 Marine Collaboration Research Forum

Capital breeding, whereby adult's build-up energy reserves at times when food is abundant in order to invest in reproduction during a time of low prey availability, is common in seasonal environments. A dependence on stored energy for reproductive investment may make capital breeders sensitive to rising temperatures during winter when activity and energy demands are typically low. The lesser sandeel, Ammodytes marinus (Raitt, 1934), is an extreme capital breeder, as gonad development is entirely dependent on stored energy whilst the fish remains buried in sand overwinter. In this laboratory study, the energetic consequences of high and low overwintering temperatures on energy allocation to reproduction in A. marinus were examined. The laboratory conditions led to growth and mass changes that were consistent with previously published field observations. Loss in wet mass was greater in the high temperature treatment, consistent with the higher metabolic cost predicted. Despite this, somatic energy loss did not differ between the two temperatures, the difference in mass being related to the lower relative gonad size in the high temperature treatment. The negative effect of temperature on reproductive investment in A. marinus highlights that although temperature can have a permissive effect on reproductive development it is limited by available energy reserves. Based on these findings it seems likely that warming will lead to a change in reproductive investment in A. marinus that mature but will not impact their overwinter survival.

Tidal energy is a renewable resource that can contribute towards meeting growing energy demands, but uncertainties remain about environmental impacts of device installation and operation. Environmental monitoring programs are used to detect and evaluate impacts caused by anthropogenic disturbances and are a mandatory requirement of project operating licenses in the United States. In the United Kingdom, consent conditions require monitoring of any adverse impacts on species of concern. While tidal turbine sites share similar physical characteristics (e.g. strong tidal flows), similarities in their biological characteristics have not been examined. To characterize the generality of biological attributes at tidal energy sites, metrics derived from acoustic backscatter describing temporal and spatial distributions of fish and macrozooplankton at Admiralty Inlet, Washington State and the Fall of Warness, Scotland were compared using t -tests, F -tests, linear regressions, spectral analysis, and extreme value analysis (EVA). EVA was used to characterize metric values that are rare but potentially associated with biological impacts, defined as relevant change as a consequence of human activity. Pelagic nekton densities were similar at both sites, as evidenced by no statistically significant difference in densities, and similar daily density patterns of pelagic nekton between sites. Biological characteristics were similar, suggesting that generic biological monitoring programs could be implemented at these two sites, which would streamline permitting, facilitate site comparison, and enable environmental impact detection associated with tidal energy deployment.

Offshore wind power generation requires large areas of sea to accommodate its activities, with increasing claims for exclusive access. As a result, pressure is placed on other established maritime uses, such as commercial fisheries. The latter sector has often been taking a back seat in the thrust to move energy production offshore, thus leading to disagreements and conflicts among the different stakeholder groups. In recognition of the latter, there has been a growing international interest in exploring the combination of multiple maritime activities in the same area (multi-use; MU), including the re-instatement of fishing activities within, or in close proximity to, offshore wind farms (OWFs). We summarise local stakeholder perspectives from two sub-national case studies (East coast of Scotland and Germany's North Sea EEZ) to scope the feasibility of combining multiple uses of the sea, such as offshore wind farms and commercial fisheries. We combined a desk-based review with 15 semi-structured qualitative interviews with key knowledge holders from both industries, regulators, and academia to aggregate key results. Drivers, barriers and resulting effects (positive and negative) for potential multi-use of fisheries and OWFs are listed and ranked (57 factors in total). Factors are of economic, social, policy, legal, and technical nature. To date, in both case study areas, the offshore wind industry has shown little interest in multi-use solutions, unless clear added value is demonstrated and no risks to their operations are involved. In contrast, the commercial fishing sector is proactive towards multi-use projects and acts as a driving force for MU developments. We provide a range of management recommendations, based on stakeholder input, to support progress towards robust decision making in relation to multi-use solutions, including required policy and regulatory framework improvements, good practice guidance, empirical studies, capacity building of stakeholders and improvements of the consultation process. Our findings represent a comprehensive depiction of the current state and key stakeholder aspirations for multi-use solutions combining fisheries and OWFs. We believe that the pathways towards robust decision making in relation to multi-use solutions suggested here are transferable to other international locations.

VC International Council for the Exploration of the Sea 2016. James J. Waggitt was funded by a NERC Case studentship supported by OpenHydro Ltd and Marine Scotland Science (NE/J500148/1). Shore-based surveys were funded by a NERC (NE/J004340/1) and a Scottish National Heritage (SNH) grant. FVCOM was funded by a NERC grant (NE/J004316/1). The bathymetry data used in hydrodynamic models (HI 1122 Sanday Sound to Westray Firth) was collected by the Maritime and Coastguard Agency (MCA) as part of the UK Civil Hydrography Programme. We wish to thank Christina Bristow, Matthew Finn and Jennifer Norris at the European Marine Energy Centre (EMEC); Ian Davies at Marine Scotland Science; Gail Davoren, Shaun Fraser, Pauline Goulet, Alex Robbins and Helen Wade for invaluable discussions; Thomas Cornulier, Alex Douglas, James Grecian and Samantha Patrick for their help with statistical analysis; and Jenny Campbell and the Cockram family for assistance during fieldwork.

Acknowledgements University of Aberdeen, UK and Bay of Bengal Large Marine Ecosystems (BOBLME) project are acknowledged for partial funding of this research.

Aim Given the paucity of data on the distribution of habitats and species for most marine species, particularly those that are rare and in need of protection, there is a need to model species distributions. Using the fan mussel, Atrina fragilis (Pennant 1777), as our case species, the aim of the study was to predict new areas of occurrence for A. fragilis, estimate the extent of potentially suitable areas, determine the proportion of these areas that are included in the recently designated nature conservation MPAs off the west of Scotland and identify possible environmental drivers in the distribution of A. fragilis. Location West coast of Scotland, UK. Methods Using a point process framework, we modelled presence-only data, including historic records. A quadrat survey employing digital still photography was conducted in areas of high and low suitability to verify the model, and subsequently, a targeted survey was undertaken in areas predicted as highly suitable by the models using towed video cameras. Results Five environmental variables were of prime importance in explaining the distribution of A. fragilis. The results from the verification survey support model performance. Atrina fragilis was found in 80% of the targeted transects undertaken. Approximately 14% of the total area predicted as suitable for A. fragilis occurred within recently designated marine protected areas (MPAs) indicating considerable potential for recolonization given suitable protection. Main conclusions The verified model suggests that limited presence-only and historical records of rare species can perform well within a SDM framework allowing the identification of further suitable areas. The prominence of bathymetric ruggedness in the models is unexpected, given the understood ecological niche for A. fragilis and pinnids in general, but is consistent with the fact that seabed topography can offer protection from fishing pressure. These results will inform restoration objectives of the MPA network.

Acknowledgements: James J. Waggitt was funded by a NERC Case studentship supported by OpenHydro Ltd and Marine Scotland Science (NE/J500148/1). Vessel-based transects were funded by a NERC (NE/J004340/1) and a Scottish National Heritage (SNH) grant. FVCOM modelling was funded by a NERC grant (NE/J004316/1). Marine Scotland Science provided time on the FRV Alba-na-Mara as part as the Marine Collaboration Research Forum (MarCRF). The bathymetry data used in hydrodynamic models (HI 1122 Sanday Sound to Westray Firth) was collected by the Maritime & Coastguard Agency (MCA) as part of the UK Civil Hydrography Programme. We wish to thank Christina Bristow, Matthew Finn and Jennifer Norris at the European Marine Energy Centre (EMEC); Marianna Chimienti, Ciaran Cronin, Tim Sykes and Stuart Thomas for performing vessel-based transects; Marine Scotland Science staff Eric Armstrong, Ian Davies, Mike Robertson, Robert Watret and Michael Stewart for their assistance; Shaun Fraser, Pauline Goulet, Alex Robbins, Helen Wade and Jared Wilson for invaluable discussions; Thomas Cornulier, Alex Douglas, James Grecian and Samantha Patrick for their help with statistical analysis; and Gavin Siriwardena, Leigh Torres, Mark Whittingham and Russell Wynn for their constructive comments on earlier versions of this manuscript. APC paid through institutional prepayment scheme

Aim Estimating environmental suitability from species distribution data is crucial in defining spatial conservation measures. To this end, species distribution models (SDMs) are commonly applied, but seldom validated by completely independent data. Here, we use data on individual tracks derived from electronic tags as an alternative means of validating SDM outputs. Location West coast of Scotland, NE Atlantic. Methods We used a binomial generalized additive model (GAM) to predict the environmental suitability for flapper skate (Dipturus cf. intermedia) in Scottish waters. The GAM modelled relative habitat usage as a function of environmental variables using presence–absence data obtained from scientific trawl surveys. Additional data obtained from electronic tags attached to six individual flapper skates were used to estimate individual tracks using a tidal-based geolocation model. Concordance between individual tracks and GAM-predicted maps of relative habitat usage (RHU) was tested by comparing predicted RHU between estimated tracks and randomly generated tracks. Results Environmental suitability for the flapper skate was driven by depth and distance from the coast in the SDM. We found high spatial concordance between the estimated tracks of the six tagged individuals and regions of high RHU predicted by the SDM. Main conclusions Integrating outputs from an independent data source allowed us to validate predictions from a species distribution model (SDM). The integration of individual- and population-level data sources increases confidence in the outputs being used to define spatial conservation measures. The information on flapper skate distribution provided by this study provides a useful framework for considering spatial conservation measures for this species.

The recent increase in data accuracy from high resolution accelerometers offers substantial potential for improved understanding and prediction of animal movements. However, current approaches used for analysing these multivariable datasets typically require existing knowledge of the behaviors of the animals to inform the behavioral classification process. These methods are thus not well‐suited for the many cases where limited knowledge of the different behaviors performed exist. Here, we introduce the use of an unsupervised learning algorithm. To illustrate the method's capability we analyse data collected using a combination of GPS and Accelerometers on two seabird species: razorbills (Alca torda) and common guillemots (Uria aalge). We applied the unsupervised learning algorithm Expectation Maximization to characterize latent behavioral states both above and below water at both individual and group level. The application of this flexible approach yielded significant new insights into the foraging strategies of the two study species, both above and below the surface of the water. In addition to general behavioral modes such as flying, floating, as well as descending and ascending phases within the water column, this approach allowed an exploration of previously unstudied and important behaviors such as searching and prey chasing/capture events. We propose that this unsupervised learning approach provides an ideal tool for the systematic analysis of such complex multivariable movement data that are increasingly being obtained with accelerometer tags across species. In particular, we recommend its application in cases where we have limited current knowledge of the behaviors performed and existing supervised learning approaches may have limited utility.

Management of the sea is increasingly complex, riddled with uncertainty and necessitates involvement from researchers across disciplines and stakeholders from multiple policy and practice sectors. This article discusses “The Cooperative Participatory Evaluation of Renewable Technologies on Ecosystem Services” (CORPORATES) research project, which developed an innovative and practical method of linking ecological processes, ecosystem services and benefits. The research was conducted in the context of licensing decisions for offshore wind farms in the North Sea (Scotland, UK). A set of linked, modular participatory processes were developed to foster cross-sector stakeholder engagement. It employed an exchange of ecological, legal, social, economic and cultural knowledge around marine ecosystem services. Workshop exercises included participatory mapping, benefit identification, and developing an understanding of linkages between ecosystem services, benefits, stakeholders’ activities and policy drivers through co-development of conceptual systems maps of the study area. The participatory exercises fostered meaningful dialogue across sectors and an ability to participate equally, despite initial differences in knowledge about ecosystem services. The development of conceptual systems maps facilitated productive discussion about trade-offs in relation to different policies. Reflective discussion identifies ways in which the developed processes could be integrated into future decision making. An assessment of the approach revealed that it operationalised a post normal science framework in terms of process oversight, multiple knowledge claims, and managing uncertainty. It developed a process that linked understanding of ecosystem functioning with the creation and implementation of policy thereby creating an ecosystem approach to marine spatial planning and licensing decisions, as required by law. This approach has extensive transferability to situations where stakeholder engagement is required to develop policy and provide feedback as part of a decision-making process. It is an engagement, outreach tool for communities and can help teach methods and processes for stakeholder engagement which enable new insights.

The generation of offshore energy is a rapidly growing sector, competing for space in an already busy seascape. This book brings together the ecological, economic, and social implications of the spatial conflict this growth entails. Covering all energy-generation types (wind, wave, tidal, oil, and gas), it explores the direct and indirect impacts the growth of offshore energy generation has on both the marine environment and the existing uses of marine space. Chapters explore main issues associated with offshore energy, such as the displacement of existing activities and the negative impacts it can have on marine species and ecosystems. Chapters also discuss how the growth of offshore energy generation presents new opportunities for collaboration and co-location with other sectors, for example, the co-location of wild-capture fisheries and wind farms. The book integrates these issues and opportunities, and demonstrates the importance of holistic marine spatial planning for optimising the location of offshore energy-generation sites. It highlights the importance of stakeholder engagement in these planning processes and the role of integrated governance, with illustrative case studies from the United States, United Kingdom, northern Europe, and the Mediterranean. It also discusses trade-off analysis and decision theory and provides a range of tools and best practices to inform future planning processes.

Despite rapid development of marine renewable energy, relatively little is known of the immediate and future impacts on the surrounding ecosystems. Quantifying the behavior and distribution of animals around marine renewable energy devices is crucial for understanding, predicting, and potentially mitigating any threats posed by these installations. The Flow and Benthic Ecology 4D (FLOWBEC) autonomous seabed platform integrated an Imagenex multibeam echosounder and a Simrad EK60 multifrequency echosounder to monitor marine life in a 120 $^{\circ}$ sector over ranges up to 50 m, seven to eight times per second. Established target detection algorithms fail within MRE sites, due to high levels of backscatter generated by the turbulent physical dynamics, limiting and biasing analysis to only periods of low current speed. This study presents novel algorithms to extract diving seabirds, fish, and fish schools from the intense backscatter caused by turbulent dynamics in flows of 4 m s $^{{-1}}$ . Filtering, detection, and tracking using a modified nearest neighbor algorithm provide robust tracking of animal behavior using the multibeam echosounder. Independent multifrequency target detection is demonstrated using the EK60 with optimally calculated thresholds, scale-sensitive filters, morphological exclusion, and frequency-response characteristics. This provides sensitive and reliable detection throughout the entire water column and at all flow speeds. Dive profiles, depth preferences, predator–prey interactions, and fish schooling behavior can be analyzed, in conjunction with the hydrodynamic impacts of marine renewable energy devices. Coregistration of targets between the acoustic instruments increases the information available, providing quantitative measures including frequency response from the EK60, and target morphology and behavioral interactions from the multibeam echosounder. The analyses draw on deployments at a tidal energy site in Scotland to compare the presence and absence of renewable energy structures across a range of physical and trophic levels over complete spring-neap tidal cycles. These results can be used to inform how animals forage in these sites and whether individuals face collision risks. This quantitative information can de-risk the licensing process and, with a greater mechanistic understanding at demonstration scales, its predictive power could reduce the monitoring required at future arrays.

Marine space is about to get even busier. Around the globe many countries are looking to use their ocean space, especially their coastal marine areas, for the extraction of renewable energy from the wind, waves and tides. This large scale use of renewable energy will help global reductions in CO2 release and ultimately reduce the risk of ecological damage due to sever climate change. However the cumulative effect of large scale developments and the range of methods to extract energy from the oceans will cause local changes in physical mixing properties and the effects will be felt locally throughout the trophic chain. How can we assess the ecological costs and benefits of these contrasting pressures on our marine systems such that our choices in where we allow the placement of large scale energy extraction is ultimately ecological beneficial? In this work we suggest that the use of statistical joint models; spatially explicit models that simultaneous explore the distributions of mobile predator and prey species such as pelagic fish and seabird and marine mammal species are a most useful tool in the calculation of the degree of overlap in these species now and in future predictions with climate change, energy extraction and both climate change and energy extraction. The contrasting degree of spatial overlap in these different scenarios will allow the estimate on the ecological costs and benefits of large scale extraction of marine renewable energy.

During the breeding season, seabirds are obligate central place foragers, and this may make them vulnerable to impacts of environmental change. An individual based model of a pair of central place foragers and their offspring has been developed for the common guillemot ( Uria aalge ). The behavioural decisions of each adult depend on the state of themselves, their partner and their chick. The behaviour of the adults and the body masses of all three are followed over the chick rearing period. The model was used to investigate how chick fledging mass, proportion of time the chick was left unattended by its parents and change in adult mass are impacted by different foraging ranges, prey abundance and calorific content. Adults in the model typically declined in mass during the chick rearing period, although none died of starvation. Provisioning parents can, to some extent, increase foraging distance from the colony without a reduction in the proportion of chicks reaching a suitable fledging mass or increasing the time they are left unattended. The foraging range at which a decline in fledging success and colony attendance occurs is influenced by changes in either one or both of, prey abundance and prey quality. Patterns produced by the model are consistent with field observations and biological knowledge. As the model outputs can give an indication of the fitness consequences of environmental changes it can be used to address theoretical ecological questions as well as to inform marine spatial management.

It is likely that there will be a substantial increase in the number of tidal stream turbines within the UK over the next decade. However, the ecological impacts upon marine top-predators, including seabirds, remain largely unknown. Although tidal stream turbines could have many direct and indirect impacts upon seabird populations, it is the risk of direct collisions between individuals and moving components that currently causes the most concern. Species such as Auks Alcidae sp., Cormorants Phalacrocorax sp. and Divers Gavia sp. almost certainly face higher risks than others. However, it is likely that they are not equally vulnerable. Part of predicting which are most vulnerable involves the estimation of spatial overlap between their foraging distributions and the location of tidal stream turbines. This paper reviews potential methods and approaches that should help to predict whether a population would: (1) exploit areas suitable for tidal stream turbines, (2) dive near tidal stream turbines within these areas, or (3) dive to depths where moving components are found? Answering these questions in a hierarchical manner (from 1 to 3) could help to predict the extent of spatial overlap for vulnerable populations. These approaches require a fundamental understanding of the mechanistic links between physical conditions, prey characteristics and foraging opportunities. Therefore, multi-disciplinary approaches incorporating methods usually associated with oceanographic and fisheries studies are needed to document physical conditions and prey characteristics over large and small spatial scales. Answering these questions also requires collaborative efforts and a strategic governance approach to collating the wide range of distributional, prey and physical datasets currently being collected.

Commercial fisheries data, collected as part of an observer programme and covering the period 1997–2014, were utilized in order to define key reproductive traits and spawning dynamics of the Patagonian toothfish Dissostichus eleginoides at South Georgia. Multi-year spawning site fidelity of D. eleginoides was revealed through the identification of previously unknown spawning hotspots. Timing of female spawning was shown to have shifted later, leading to a shorter spawning duration. A decrease in length and mass of female and male spawning fish and a reduced number of large spawning fish was found, evidence of a change in size structure of spawning D. eleginoides. During the study period fewer later maturity stage females (including spawning stage) were observed in conjunction with increased proportions of early stage female D. eleginoides. The findings are discussed in the context of reproductive success, with consideration of the possible effects such spawning characteristics and behaviours may have on egg and larval survival. This work presents the first long-term assessment of D. eleginoides spawning dynamics at South Georgia and provides valuable knowledge for both the ecology of the species and for future fisheries management of this commercially important species.

The Celtic Sea is a diverse fishing ground that supports important commercial fisheries for a range of demersal fish, large and small-bodied pelagic fish and a variety of cephalopods and other shellfish. A regional overview of the main commercial fish stocks of the Celtic Sea and of the fish that occur in the vicinity of Jones Bank are provided through analyses of landings data from English and Welsh vessels, and from scientific trawl surveys. Dedicated smaller scale sampling via trawl surveys combined with baited cameras on and around the Jones Bank were also analysed to investigate the importance of sandbank habitats with attention paid to the differences in the species occurring on the top of the bank in comparison to adjacent off-bank habitats. Official landing statistics for UK (English and Welsh) vessels indicated that the predominant commercial demersal species in ICES Divisions VIIg,h (in terms of quantities landed) were anglerfish, megrim, pollack and skates (Rajidae). There were, however, regional differences in the distribution of fish and fisheries, and the area surrounding Jones Bank (ICES Rectangles 28E1 and 28E2) supports fisheries for megrim, anglerfish, skates, hake, ling and turbot, with otter trawl, gillnet and beam trawl the main gears used. Recent survey data collected with GOV (Grande Ouverture Verticale) trawl from the Celtic Sea (ICES Divisions VIIe-h, 2007–2010) were used to highlight the broad scale distribution of the main fish assemblages in the Celtic Sea. Analyses of the fish and cephalopod catches from these surveys indicated that there were four broad assemblages in the area, including (i) a region around the Cornwall (which will also be partly influenced by the necessity to use rockhopper ground gear on these rough grounds), (ii) the shallower regions of the north-western Celtic Sea (including parts of the Bristol Channel), (iii) the deeper parts of the outer shelf and (iv) the central Celtic Sea. These data also provided information on the ichthyofauna of the Jones Bank. Further site-specific data for bank and off-bank habitats were collected during dedicated surveys on the Jones Bank in 2008 using commercial trawlers and baited camera deployments. Twenty-three species were recorded on the top of the bank, where horse mackerel, haddock and boarfish were the most abundant species; 18 species were found along the slope of the bank (with blue whiting, poor cod, hake and horse mackerel predominant) and 18 species observed off the bank (where catches were dominated by blue whiting, poor cod and hake). The differences between camera and trawls were important with cameras only picking up 28% of the species seen in the trawls. However both camera and trawl results suggest that some species are very habitat specific, with species such as haddock only observed on the top of the bank, whilst Nephrops norvegicus was abundant on the flat areas off the bank but was infrequent on the top of the bank. These results suggest that future surveys of offshore sandbank habitats should stratify sampling more specifically to deal with smaller scale features that may play an important role in providing a greater range of habitats than just their relative size would suggest.

Fishing vessel position data from the Vessel Monitoring System (VMS) were used to investigate fishing activity in the Celtic Sea, a seasonally-stratifying, temperate region on the shelf of northwest Europe. The spatial pattern of fishing showed that three main areas are targeted: (1) the Celtic Deep (an area of deeper water with fine sediments), (2) the shelf edge, and (3) an area covering several large seabed banks in the central Celtic Sea. Data from each of these regions were analysed to examine the contrasting seasonality of fishing activity, and to highlight where the spring-neap tidal cycle appears to be important to fishing. The oceanographic characteristics of the Celtic Sea were considered alongside the distribution and timing of fishing, illustrating likely contrasts in the underlying environmental drivers of the different fished regions. In the central Celtic Sea, fishing mainly occurred during the stratified period between April and August. Based on evidence provided in other papers of this Special Issue, we suggest that the fishing in this area is supported by (1) a broad increase in primary production caused by lee-waves generated by seabed banks around spring tides driving large supplies of nutrients into the photic zone, and (2) greater concentrations of zooplankton within the region influenced by the seabed banks and elevated primary production. In contrast, while the shelf edge is a site of elevated surface chlorophyll, previous work has suggested that the periodic mixing generated by an internal tide at the shelf edge alters the size–structure of the phytoplankton community which fish larvae from the spawning stocks along the shelf edge are able to exploit. The fishery for Nephrops norvegicus in the Celtic Deep was the only one to show a significant spring-neap cycle, possibly linked to Nephrops foraging outside their burrows less during spring tides. More tentatively, the fishery for Nephrops correlated most strongly with a localised shift in the tidal current polarisation, suggesting that the muddy seabed required by Nephrops is controlled by rotational constraints on the extent of the bottom boundary layer.

The epibenthic assemblages in the Celtic Sea are described from the catches from 2 m beam trawl surveys undertaken from 2000 to 2009. During this period 154 samples were collected. The most ubiquitous species in the study area were the natantid shrimps Processa spp. and Crangon allmanni, the hermit crabs Pagurus prideaux and Anapagurus laevis, sand star Astropecten irregularis and spotted dragonet Callionymus maculatus. Multivariate community analyses indicated that catches (numbers per tow) were distributed across six assemblages, two of which were predominant in the study area. Most catches were attributed to either a ‘shelf edge assemblage‘, which was widespread in deeper waters (114–423 m water depth) or an ‘outer shelf assemblage’ that occurred across much of the Celtic Sea north of 49°N in waters 49–175 m deep. The dominant species along the edge of the continental shelf were the hormathid anemome Actinauge richardi, sea spider Pycnogonum littorale (which associated with A. richardi), Devonshire cup coral Caryophyllia smithii and the swimming crab Macropipus tuberculatus. The dominant species in the outer shelf assemblage included P. prideaux, C. allmanni, A. laevis and common starfish Asterias rubens. Stations closer to shore were relatively distinct and catches in this ‘inner shelf assemblage’ were composed primarily of an inshore fauna (e.g. Ophiura ophiura, C. allmanni and Liocarcinus holsatus). Stations in the southern part of the survey grid were also relatively distinct (‘southern Celtic Sea assemblage’), and several large echinoderms (Porania pulvillus, Stichastrella rosea and Anseropoda placenta) dominated at these sites. Three of the deepest stations were also relatively distinct, as were a group of stations in the muddy habitat of the Celtic Deep and comparable grounds elsewhere in the region, where Nucula sulcata and Alpheus glaber were characteristic. Catches on the shallower parts of the Jones Bank (and on another bank in the region) were dominated by the anemone Paraphellia expansa, with off-bank sites comprising a greater number of species. In contrast to beam trawl sampling, baited camera observations on the Jones Bank showed a greater richness of species on the shallower part of the bank, and provided information on the nocturnal feeding behaviour of scavenging isopods.

As we begin to manage our oceans in much more spatial detail we must understand a great deal more about oceanographic habitat preferences of marine mobile top predators. In this unique field study we test a hypothesis on the mechanisms defining mobile predator foraging habitat characteristics by comparing temporally and spatially detailed bio-physical oceanographic data from contrasting topographical locations. We contrast the foraging locations of two very different seabird species, gannets and storm petrels, by repeatedly sampling a bank and a nearby flat area over daily tidal cycles during spring and neap tides. The results suggest that storm petrels are linked to foraging in specific locations where internal waves are produced, which is mainly on banks. These locations can also include the presence of high biomass of chlorophyll. In contrast, the location where more gannets are foraging is significantly influenced by temporal variables with higher densities of foraging birds much more likely during the neap tide than times of spring tide. The foraging times of both species was influenced by differences between the vertical layers of the water column above and below the thermocline; via either vertical shear of horizontal currents or absolute differences in speed between layers. Higher densities of foraging gannets were significantly more likely to be found at ebb tides in both bank and flat regions however over the bank, the density of foraging gannets was higher when the differences in speed between the layers were at a maximum. Both gannets and storm petrels appear to be more likely to forage when wind direction is opposed to tidal direction. This detailed understanding links foraging behaviour to predictable spatial and temporal bio-physical vertical characteristics and thus can be immediately used to explain variance and increase certainty in past abundance and distributional surveys. These results also illuminate the types of variables that should be considered when assessing potential changes to the distribution and characteristics of habitats from increased anthropogenic disturbances such as large scale offshore wind, wave and tidal renewable deployments.